Original Paper
Brain Behav Evol 2021;96:1–12
A Farewell to the Encephalization
Quotient: A New Brain Size Measure for
Comparative Primate Cognition
Carel P. van Schaik
a, b
Zegni Triki
c, d
Redouan Bshary
c
Sandra A. Heldstab
a
a
Department of Anthropology and Anthropological Museum, University of Zurich, Zurich, Switzerland;
b
Department of Evolutionary Biology and Environmental Science, University of Zurich, Zurich, Switzerland;
c
Behavioral Ecology Laboratory, Faculty of Science, University of Neuchâtel, Neuchâtel, Switzerland;
d
Institute of
Zoology, Stockholm University, Stockholm, Sweden
Received: February 17, 2021
Accepted: May 2, 2021
Published online: July 9, 2021
Correspondence to:
Carel P. van Schaik, vschaik @aim.uzh.ch
© 2021 The Author(s)
Published by S. Karger AG, Basel
karger@karger.com
www.karger.com/bbe
DOI: 10.1159/000517013
Keywords
Encephalization quotient · Cognitive equivalence ·
Intelligence · Mammals · Hominins
Abstract
Both absolute and relative brain sizes vary greatly among
and within the major vertebrate lineages. Scientists have
long debated how larger brains in primates and hominins
translate into greater cognitive performance, and in particu-
lar how to control for the relationship between the noncog-
nitive functions of the brain and body size. One solution to
this problem is to establish the slope of cognitive equiva-
lence, i.e., the line connecting organisms with an identical
bauplan but different body sizes. The original approach to
estimate this slope through intraspecific regressions was
abandoned after it became clear that it generated slopes
that were too low by an unknown margin due to estimation
error. Here, we revisit this method. We control for the error
problem by focusing on highly dimorphic primate species
with large sample sizes and fitting a line through the mean
values for adult females and males. We obtain the best esti-
mate for the slope of circa 0.27, a value much lower than
those constructed using all mammal species and close to the
value expected based on the genetic correlation between
brain size and body size. We also find that the estimate of
cognitive brain size based on cognitive equivalence fits em-
pirical cognitive studies better than the encephalization
quotient, which should therefore be avoided in future stud-
ies on primates and presumably mammals and birds in gen-
eral. The use of residuals from the line of cognitive equiva-
lence may change conclusions concerning the cognitive
abilities of extant and extinct primate species, including
hominins. © 2021 The Author(s)
Published by S. Karger AG, Basel
Introduction
Although recent ecological approaches to comparative
cognition have focused on linking performance in spe-
cific cognitive tasks to specific brain regions [e.g., Healy
and Krebs, 1996], traditionally comparative cognition has
relied on a presumed link between some summary mea-
sure of cognitive performance and total brain size [Jeri-
son, 1973]. Scholars have therefore long been searching
for a neuroanatomical measure of overall cognitive abil-
ity, both to compare living species and to estimate the
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